Hyperossification in the vertebral column of Devonian placoderm fishes (Arthrodira)

ABSTRACT

Skeletal bone enlargement (hyperossification) was thought to only occur in endochondral and dermal bones (e.g., endoskeletons of marine tetrapods and dermal bones of certain jawless vertebrates, placoderms, and teleost fishes). However, in some arthrodiran placoderms (basal jawed vertebrates), i.e., Millerosteus minor, Compagopiscis croucheri, Eastmanosteus calliaspis, several types of bone enlargement also occur within the endoskeleton, affecting the vertebral column. Significantly, placoderm endoskeletal bone is thought to be thin-walled, ossifying only in the fibrous layer surrounding a cartilage precursor (historically called perichondral bone), rather than endochondrally/dermally. Hyperossification differs among these three species, revealing a range of internal and external bone morphologies undescribed for the placoderm endoskeleton. Thus, neural arches of M. minor are swollen in external appearance, in cross-section showing considerable deposition of layered, compact bone. In contrast, the arches of E. calliaspis are unswollen externally, with the neural spine cortex composed of the thin perichondral bone expected for placoderms. Histologically, though, the arch comprises numerous layers of perichondral bone. In C. croucheri, as in E. calliaspis, the spine and arch are unswollen externally, but the perichondral bone is thickened and compact, more similar to the condition in M. minor. Vertebral hyperossification in these taxa differs considerably in the degree and mode of cortical tissue thickening, via addition of tissue to the bone’s external (periosteal) and/or internal (endosteal) surfaces. Hyperossifcation in these arthrodires demonstrates that increases in endoskeletal bone mass are not restricted to crown-group gnathostomes (Chondrichthyes + Actinopterygii), representing a first step in the evolution of this process, involving modifications to cortical bone layers.

ACKNOWLEDGMENTS

The authors thank S. Walsh for much-needed help and guidance concerning the NMS collections, E. Bernard (Department of Earth Sciences) for access to the NHM fossil fish collections, B. Clark and V. Fernandez (Image and Analysis Centre, NHM) for access to the CT scanners and support, H. Armer (Sackler Imaging Suite, NHM), for teaching S.v.M. how to use the various microscopes and cameras in the NHM and to become much better at photography for preparation of Millerosteus minor specimens; and M. Newman for very helpful advice and guidance on field work.